EFFECT OF SILICON FERTILIZATION ON THE EXPRESSION OF ISOENZYMES,
THE GERMINATION AND SEED YIELD OF WHEAT
Lizandro Ciciliano Tavares1, André Oliveira de Mendonça1, André Pich Brunes1, Gizele
Ingrid Gadotti 2, Géri Eduardo Meneghello3
1-Pós-Graduando pelo Programa de Pós-Graduação em Ciência e Tecnologia de
Sementes – UFPel/FAEM. Campus Universitário – Caixa Postal 354 – CEP 96001-970,
Pelotas - RS. [email protected].
2-Professora adjunta, Centro de Engenharias, Universidade Federal de Pelotas.
3- Engenheiro agrônomo, doutor, Programa de Pós-Graduação em Ciência e Tecnologia
de Sementes – UFPel/FAEM. Campus Universitário – Caixa Postal 354 – CEP 96001970, Pelotas - RS.
Recebido em: 30/09/2013 – Aprovado em: 08/11/2013 – Publicado em: 01/12/2013
ABSTRACT
This study evaluated the effects of silicon fertilization on expression of isoenzymes,
germination and seed yield of wheat. The experimental design was completely
randomized with four replications. The treatments consisted of combinations of two
sources of silicon and six levels in a factorial AxB, totaling 12 treatments with four
replications. The isoenzyme patterns were studied: esterase, glutamate oxaloacetate
transaminase, malate dehydrogenase and peroxidase. After harvesting, it was
evaluated: number of seeds per plant, thousand seed weight, hectoliter weight and
seeds yield. It is concluded that silicon fertilization, with both sources, does not affect the
germination of wheat seeds, but further increase in productivity up to a dose of 1720.84
kg ha-1. The source of rice husk ash promotes greater number of ears per plant and
higher seed yield per plant than the kaolin. Electrophoretic variation of isozymes is
associated with the supply of silicon in the soil in wheat plants.
KEYWORDS: Triticum aestivum L., silicon, isoenzyme, electrophoresis, productivity.
EFEITO DA ADUBAÇÃO SILICATADA NA EXPRESSÃO ISOENZIMÁTICA, NA
GERMINAÇÃO E NO RENDIMENTO DE SEMENTES DE TRIGO
RESUMO
Objetivou-se com o presente trabalho avaliar os efeitos da adubação silicatada na
expressão isoenzimática, na germinação e no rendimento em sementes de trigo. O
delineamento experimental foi inteiramente casualizado, com quatro repetições. Os
tratamentos consistiram em combinações de duas fontes de silício, em esquema fatorial
AxB (Fator A: Caulim e cinza de casca de arroz, Fator B: níveis de 0, 500, 1000, 1500,
2000 e 2500 kg ha-1), totalizando 12 tratamentos, com quatro repetições. Os padrões
isoenzimáticos estudados foram esterase (EST, EC 3.1.1.1), glutamato oxaloacetato
transaminase (GOT, EC 2.6.1.1), malato desidrogenase (MDH, EC 1.1.1.37) e
peroxidase (PO, EC 1.11.1.7). Após a colheita avaliaram-se as seguintes variáveis:
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número de sementes por planta, peso de mil sementes, peso hectolítrico e rendimento
de sementes. Concluiu-se que a adubação silicatada, com ambas as fontes, não afetam
a germinação de sementes de trigo, porém promovem aumento na produtividade, até a
dose de 1720,84 kg ha-1. A fonte cinza de casca de arroz promove maior número de
espigas por planta e maior rendimento de sementes por planta do que o caulim.
Variações eletroforéticas de isoenzimas estão associadas ao fornecimento de silício via
solo em plantas de trigo, com exceção da malato desidrogenase (MDH, EC 1.1.1.37).
PALAVRAS-CHAVE: Triticum aestivum L., silício, isoenzimas, eletroforese,
produtividade.
INTRODUCTION
The increasing of production and productivity of major crops, among them wheat,
rice, soybeans, corn and beans are crucial to be met the demands of food caused by
population growth. Given this, there are aspects that are essential to achieve this goal,
as the use of high quality seeds, plant protection and deployment of technologies for
increasing crop productivity, emphasizing the use of nutrients, especially those classified
as essential, and the beneficial species of the family Poaceae, such as silicon (FAO,
2002).
The incidence of diseases has been a limiting factor in the full expression of yield
potential of wheat in Brazil (BACALTCHUK et al., 2006). Among the research for
improvement of agricultural production, have focused on seed technology (VIGANO et
al., 2010; TOLEDO et al., 2011). According to COPELAND & MCDONALD (2001), plant
nutrition is one of the factors that may influence seed vigor. SÁ (1994) emphasizes that
well-nourished plants can produce more seeds with better physiological quality, because
they can become more tolerant to adverse weather.
Among the nutrients beneficial to plants is silicon (Si), which provides an increase
in growth, reducing the incidence of pathogens, improvement of leaf architecture,
reduction in transpiration rate, increasing in photosynthetic rate and improves the quality
of physiological seeds, providing an increase in yield (LIMA-FILHO, 2004; LIMA-FILHO,
2005). Several authors have reported the importance of silicon for increased resistance
and reduced the incidence of fungal diseases, increase in soil fertility, and reduced
pesticide costs and environmental impact (DATNOFF et al., 2007; LIMA-FILHO & TSAI,
2007, VIEIRA et al., 2011).
Plants are quite different in their ability to absorb the Si, which is absorbed in the
acid form mono silica (H4SiO4) passively or actively, by transporters specific membrane
for this purpose. The transport and accumulation of silicon can also be regulated by an
active process that is triggered by stimulation of protection against diseases, pests and
other stress conditions (PERRY, 2007), which leaves the main body of silicon deposit. In
plants, approximately 99% of silicon is deposited in the form of silica and less than 1% is
ionic or colloidal fraction, a soluble form. Deposits of silica in plants will result in
increased production and product quality (BLAICH & GRUNDHÖFER, 1998).
To assess the quality of plants has been used in electrophoresis study of
enzymes, or isozymes with respect not only to changes in quality, but also in genetic and
biochemical regulations (ISTA, 1992), and analyzing proteins and nucleic acids, and a
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versatile technique, relatively simple, quick and very informative power (ACQUAAD,
1992).
The isoenzymes are products of gene expression and highly influenced by the
environment and therefore the management, because the genes controlling their
expression are manifested in certain developmental stages and in specific organs and
tissues, or under a given stimulus (RAMIREZ et al., 1991). In simple terms, the
variations may be considered isoenzymes of a particular enzyme in an organism which
exhibit the same substrate specificity. According to MALONE et al. (2006), the intensity
of the bands and isoenzyme profiles are specific to a particular part of the plant tissue
and developmental stage. It has been as important isozymes for checking the quality of
plants and seed: esterases (EST), glutamate oxaloacetate transaminase (GOT), malate
dehydrogenase (MDH) and peroxidase (PO) (SPINOLA et al., 2000; MENEZES et al.,
2004; SANTOS et al., 2005; VIEIRA et al., 2009).
In this context, the aim with this study to evaluate the effect of silicon fertilization
on the expression of isozymes, the germination and seed yield of wheat.
MATERIAL AND METHODS
The experiment was conducted in Seed Laboratory and Bio-Sementes in Plant
Science Department of Agronomy College “Eliseu Maciel”, Federal University of Pelotas
(UFPel/FAEM), in crop 2011 with FUNDACEP Horizonte wheat cultivar.
The wheat plants were grown in pots of 15 liters, 10 seeds were sown per pot,
only the remaining four plants per pot (high vigor), filled with sieved soil, collected from
the A1 horizon of a Haplic Planossolo Solodic Eutric (STRECK et al., 2008), belonging
to map unit Pelotas. The fertilization was performed according to the results of soil
analysis and recommendations of the CHEMISTRY AND SOIL FERTILITY - RS / SC
(2004). It was used only nitrogen, phosphorus and potassium, applied 14 days before
seeding and liming was held thirty days before sowing. After seeding the buckets were
irrigated daily maintaining the soil at field capacity.
The treatments consisted of combinations of two basic sources of silicon in a
factorial AxB (Factor A: Kaolin and rice husk ash, Factor B: levels of 0, 500, 1000, 1500,
2000 and 2500 kg ha-1), totaling 12 treatments with four replications. The application of
silicon was carried out with the seeding.
Manual harvesting was done at the stadium in which 2/3 spikelet color was pale
yellow or cream, characterizing the physiological maturity. After evaluating the following
variables: number of spikes (NS) and number of seeds per plant (NSP): performed by
manual counting of the spikes and seeds in each experimental unit. Yield (Y) was
obtained by weighing the harvested seed, and the moisture adjusted to 13%. Hectoliter
weight (HW) performed with four replicates and specific scale, with 1 liter of seeds, the
results were expressed as kg hL-1. Thousand seed weight (TSW) - were used eight
replicates of 100 seeds. For weighing calculate the mean, standard deviation and
coefficient of variation. All plots showed a coefficient of variation less than four, so we
multiplied the average by 10, and thus gave the thousand seed weight (BRASIL, 2009).
The seed quality was evaluated by tests: Germination (G) - performed with four
replicates of 50 seeds for each treatment, germination paper substrate ("germitest"),
previously moistened with distilled water, using the ratio 2.5 times the weight of dry
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paper, and maintained at 20 ° C. The evaluations we re performed according to the Rules
for Seed Analysis (BRASIL, 2009) to eight days after sowing.
The isoenzyme patterns were studied: esterase (EST, EC 3.1.1.1), glutamate
oxaloacetate transaminase (GOT, EC 2.6.1.1), malate dehydrogenase (MDH, EC
1.1.1.37) and peroxidase (PO, EC 1.11.1.7). For this, a plant was collected at 30 days
after emergence (shoot) per experimental unit at random and macerated in a porcelain
mortar in each treatment. From each sample, 200 mg of mash were placed in Eppendorf
tube together with the extraction solution (Tris-0.1 M citrate buffer pH 8.3 + 0.1 M lithium
borate pH 8.3 + 0.15% 2 -mercaptoethanol) 1:2 (w / v). Electrophoresis was performed
in 7% polyacrylamide gels, placing 20µl of each sample in holes made with the aid of a
comb acrylic. The enzyme standards were analyzed according to the methodology
described by SCANDÁLIOS (1969) and ALFENAS (1998). Gels were placed in stainless
electrophoretic vertical kept at room temperature. The electrophoretic migration was
performed using a potential difference of 10V.cm-1, until the front end formed by the
bromophenol blue reached the bottom edge of the gel. Gels were developed for the
above enzyme systems as (SCANDÁLIOS, 1969; ALFENAS, 1998) and fixed in 10%
glycerol solution.
Interpretation of results was based on visual analysis of the gels electrophoresis,
considering the presence / absence, as well as the intensity of each of the
electrophoretic bands. Since the germination data, number of spikes, number of grains
per plant, yield, test weight and grain weight were subjected to analysis of variance,
which was performed comparing the means for the qualitative factor by Tukey test and
the factor quantitative proceeded polynomial regression at 5% probability. When the
interaction was significant developments were carried out due, if not meaningful to
compare means and polynomial regression was performed with the average levels of
the respective factor. For statistical analysis we used the Statistical Analysis System
version 1.0 Winstat (MACHADO & CONCEIÇÃO, 2003).
RESULTS AND DISCUSSION
There were no significant differences in the germination of wheat seeds from
plants fertilized with kaolin and rice husk ash in the soil (Table 1). Germination at all
doses analyzed was above 80%, with the standards required for seed, where the
minimum required is 80% (BRASIL, 2005). These results run counter to those obtained
by TOLEDO et al. (2011), which used doses of 0, 150, 300 and 450 mg dm-3 silicon as
potassium silicate in oat seeds, and had greater germination at doses of 300 and 450
mg dm-3, similar results were found in the search for VIEIRA et al. (2011), where there is
a tendency of increase in germination of rice seeds with application of calcium silicate to
the dose of 1600 kg ha-1.
Data of germination, number of spikes per plant, number of seeds per plant,
seed yield per plant, hectoliter weight and thousand seed weight are in Table 1, below.
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TABLE 1. Germination (G), number of spikes per plant (NS), number of seeds per plant
(NSP), seed yield per plant (Y), hectoliter weight (HW) and thousand seed
weight (TSW) from wheat fertilization with Kaolin and rice husk ash (RHA) in
the soil. Capão do Leão/RS, 2011
Fonte
Dose Kaolin RHA Kaolin RHA Kaolin RHA Kaolin RHA
(kg ha-1)
G (%)
NS
NSP
Y (g)
0
98
99
28
29
203
204 42,4 48,3
500
100
100
24
28
219
215 42,7 46,5
1000
98
99
25
30
228
226 45,4 48,4
1500 100
100
26
30
239
236 46,8 50,2
2000
99
100
26
27
236
245 47,5 48,9
2500 100
99
25
24
220
238 45,2 47,8
Avg 99 A 100 A 26 B 28 A 224 A 227 A 45.0 B 48.4 A
V.C.(%)
1,2
10,8
9,8
7,6
Kaolin RHA
HP (kg. hL-1)
78,0 78,9
78,4 78,8
79,0 78,9
79,1 79,3
79,2 78,8
79,4 78,6
78.9 A 78.9 A
0,5
Kaolin RHA
TSW (g)
43,7 43,1
43,1 43,2
43,0 42,7
42,8 42,6
42,7 42,8
42,2 42,7
42,9 A 42,9 A
1,8
* Means followed by same capital letter in the line in each response variable, do not
differ by Tukey test at 5% probability.
The source of silicon from rice husk ash (RHA) showed a significant increase in
the number of spikes and yield to source Kaolin (Table 1). However, the number of
seeds per plant, hectoliter weight and thousand seed weight showed no statistical
difference between the two sources studied. For all these variables there was no
interaction between doses and sources. The results agree with those obtained by LIMAFILHO & TSAI (2007), where testing different doses of silicon in three wheat cultivars
and two of oats, observed a significant increase in the number of spikes in the three
wheat cultivars studied, obtaining significant increase in productivity in both species, but
more marked in wheat, where the increase in seed production reached 100%. Similar
data were found by SINGH et al. (2005) who studied doses and application times of
silicon in two consecutive harvests of rice, where the silicon fertilization increased the
height, dry matter production, number of panicles per square meter and productivity.
For the variable number of seeds per plant (Figure 1A), there was an increase to
the point of maximum efficiency at a dose of 1808.4 kg ha-1 of silicon. The increase seed
production may be related to the highest photosynthetic as silicon to the architecture of
the plant (DEREN et al., 1994), resulting in a smaller opening angle leaf capitation
allowing more light energy (YOSHIDA et al., 1969), increasing productivity. The data
corroborate the results found by LIANG et al. (1994), which found that implementation of
silicon in the soil in four consecutive years, increased productivity of rice and wheat from
4.6 to 20.7% and from 4.1 to 9.3% respectively. ZAGO et al. (2010), using soybean foliar
application, in the doses 0, 2, 4, 6 and 8 L ha-1 had no significant difference in number of
seed per plant.
To assess the yield (Figure 1B), the sources used to promote increase the dose
of 1720.84 kg ha-1 silicon, which is its point of maximum. BARBOSA-FILHO et al.
(2001), working with three rice cultivars, found that the seed yield increased significantly
and linearly with increasing doses of application of silicon, in the order of 0.002% for
each mg SiO2.kg -1 soil. However, these data disagree with those obtained by
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CARVALHO-PUPATTO et al., (2004), working with the rice cultivar IAC-202 in field
conditions, did not achieve significant results.
1A
1B
1C
1D
FIGURE 1. Average of the sources of silicon applied to soil in the number of seeds
per plant (1A), seed yield per plant (1B), hectoliter weight (1C) and
thousand seed weight (1D) of wheat. Capão do Leão-RS, 2011.
As for hectoliter weight (Figure 1C), we observed an increase in this variable to its
maximum, which was the dose of 1784.8 kg ha-1 with subsequent reduction. The data
are consistent with those found by REIS et al. (2008), who found an increase in
hectoliter weight in rice cultivars IAC 201 and IAC 202 with increasing doses of silicon
being, however this increase showed a linear increase. However the data presented by
ORIOLI-JÚNIOR et al. (2008), working with the rice cultivars IAC 24 and IAC 370, differ
from those obtained by this project since no significant differences for these variables as
the silicon levels used.
Unlike the increase appears to other variables, the thousand seed weight was
reduced in a linear tendence of 3x10-4 g per unit dose increase of silicon (Figure 1D).
This effect may be due to a possible toxicity of the doses. The results disagree with
those found by LIMA FILHO & TSAI (2007), who found positive results for the weight of
seeds up to 25%, 44% and 35% for wheat cultivars BR18, BR40 and IPR85,
respectively. JUNIOR ORIOLI et al. (2008) using wheat cultivars IAC 24 and IAC 370
showed no influence on the silicon levels evaluated for thousand seed weight.
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For electrophoretic patterns are shown in Figures 2 and 3. Notes to the source
kaolin was reduced in intensity in the band of esterase (EST, EC 3.1.1.1) at a dose of
2500 kg ha-1 (Figure 2A), but for the source of rice husk ash, there was a reduction
significant expression of this enzyme in doses 2000 and 2500 kg ha-1. It is inferred that,
because this enzyme is responsible for both the hydrolysis of esters, and lipid
metabolism, changes in the patterns of the same evidence the occurrence of
deterioration (SANTOS et al., 2004; TUNES et al., 2010), and lipid peroxidation
according to BASAVARAJAPPA et al., (1991), an event associated with degenerative
process membranes. The data found to yield confirm the effect of seed decay, given that
the use of a silicon source rice hull ash in the soil, showed lower expression of the
esterase.
A
B
FIGURE 2. Electrophoresis patterns obtained with the EST (2A) and GOT (2B)
isoenzyme system in wheat plants subjected to different sources and
levels of silicon in the soil.
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For the standard isoenzyme glutamate oxaloacetate transaminase (GOT, EC
2.6.1.1), shown in Figure 2B, there is no difference in the intensity of the bands in both
sources. This is an enzyme that participates in the process of degradation and synthesis
of amino acids (CONN & STUMPF, 1980), presenting an important role in seed
germination, agreeing with the results of this study, which showed no difference in
germination rates, resulting in a high yield. This enzyme is responsible for the oxidation
of amino acids, providing power to the Krebs Cycle or reduced α-ketoglutarate for the
synthesis of new amino acids, such as power supply to the developing embryo (VIEIRA
et al., 2009). In this function is directly involved in the metabolism of N, it is possible for
variations occurring as happens the synthesis and degradation of amino acids during
the germination process. Undoubtedly, the enzyme GOT has a fundamental role in
protein metabolism, not only during germination, but throughout the life cycle of the
plant.
A
B
FIGURE 3. Electrophoresis patterns obtained with the system malate dehydrogenase MDH (3A) and
PO (3B) in wheat plants subjected to different sources and levels of silicon in the soil.
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Analyzing the system for the enzyme malate dehydrogenase (MDH, EC 1.1.1.37)
(Figure 3A), no difference is observed in the expression. The malate dehydrogenase
enzyme catalyzes the conversion of malate to oxaloacetate, and the important function
of NADH for the production of the Krebs cycle and generation of oxaloacetate to
biossínteses amino acid. According to SATTERS et al. (1994), because it is an important
enzyme in the cell breathing process, the increase in activity may be due to increased
expression of this enzyme in various cellular compartments and / or the induction of
enzyme activity expressed by a higher intensity of bands, confirming the data positives
obtained in the yield, since an increase in rise of the mean breathing deteriorating
process.
For the electrophoresis pattern obtained in the peroxidase (PO, EC 1.11.1.7)
(Figure 3B) shows an increase in intensity of bands for the dose of 2000 kg ha-1 Kaolin
the source. Peroxidases are proteins of approximately 50 kDa which are present as
multiple isoenzymes in plant tissues (JEBARA et al., 2005). Are associated with
physiological and biochemical processes such as growth, cell formation, fruit
development, ethylene biosynthesis and response to various stresses (MATAMOROS et
al., 2003). They participate in processes related to cell wall, such as oxidation of phenols
and lignification of plant cells during host defense response to pathogens (DATTA &
MUTHUKRISHNAN, 1999). The increased expression of this enzyme as increases the
amount of silicon corroborates the data obtained in this work, where the average of the
sources had the highest yield at a dose of 1720.84 kg ha-1 Si, implying that a good
sanity for plants with better performance.
The use of silicon source for rice husk ash is promising, because it proved
superior to kaolin analyzed when the number of spikes per plant and yield, besides
being an alternative source and more affordable cost. It was further observed that silicon
increased the number of seeds per plant, yield and hectoliter weight with increasing
dose.
As for the four different enzyme systems studied, the enzyme esterase (EST, EC
3.1.1.1), glutamate oxaloacetate dehydrogenase (GOT, EC 2.6.1.1) and peroxidase (PO,
EC 1.11.1.7), proved to be as promising biochemical markers for assessing the degree
of deterioration of wheat plants. It was detected in the electrophoretic profiles of plants
subjected to different silicon levels, showing that this may lead to lower levels
deteriorative and induction of plant resistance, resulting thus to an increase in
productivity.
The results of this study suggest that, depending on the enzyme system used,
there is a differentiation of proteins. As a result, the combined analysis of several
isozymes systems and recommend modifications to allow verification that occur inside
the plant when subjected to some kind of treatment that influences productivity.
CONCLUSIONS
Silicon fertilization, with both sources, does not affect the germination of wheat
seeds, but further increase in productivity up to a dose of 1720.84 kg ha-1. The source of
rice husk ash promotes greater number of spikes per plant and higher seed yield per
plant than the kaolin.
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Electrophoretic variation of isozymes are associated with the supply of silicon in
the soil in wheat plants, with the exception of malate dehydrogenase (MDH, EC
1.1.1.37).
REFERENCES
ACQUAAD, G. Practical protein electrophoresis for genetic research. Portland:
Discorides Press, 1992, 131 p.
ALFENAS, A. C. Eletroforese de isoenzimas e proteínas afins. Viçosa: UFV, 1998,
574 p.
BACALTCHUK, B.; CHAVES, M. S.; LIMA, M. I. P. M.; COSTAMILAN, L. M.; MACIEL, J.
L. N.; SALVADORI, J. R.; GAMBATTO, A. Características e cuidados com algumas
doenças
de
trigo.
Passo
Fundo:
Embrapa
–
trigo,
2006.
http://www.cnpt.embrapa.br/biblio/do/p_do64.pdf
BARBOSA-FILHO, M. P.; SNYDER, G. H.; FAGERIA, N. K.; DATNOFF, L. E.; SILVA, O.
F. Silicato de cálcio como fonte de silício para o arroz de sequeiro. R. Bras. Ci. Solo, v.
25, p. 325-30, 2001. http://www.malavolta.com.br/pdf/502.pdf
BASAVARAJAPPA, B. S.; SHETTY, H. S.; PRAKASH, H. S. Membrane deterioration and
other biochemical changes, associated with accelerated ageing of maize seeds. Seed
Science
and
Technology,
v.
19,
n.
2,
p.
279-286,
1991.
http://www.refdoc.fr/Detailnotice?cpsidt=5209145&traduire=en
BLAICH, R.; GRUNDHÖFER, H. Silicate incrusts induced by powdery mildew in cell
walls of different plant species. Zeitschift fur Pflanzenkrankheiten und Pflanzenschutz,
p.114-120, 1998. http://cat.inist.fr/?aModele=afficheN&cpsidt=2308876
BRASIL. Ministério da Agricultura, Pecuária e Abastecimento. 2005. Padrões para
produção e comercialização de sementes de trigo e de trigo duro. Anexo XII
(Instrução
Normativa
MAPA,
n°
25
de
16/12/2005).
http://www.google.com.br/url?sa=t&rct=j&q=&esrc=s&source=web&cd=4&ved=0CFgQFj
AD&url=http%3A%2F%2Fagrosem.agr.br%2Fsite%2Fdownload%2FProducao%2520%2520In%252025.doc&ei=HX_PT7fpCJOJ6gHqoKyJDA&usg=AFQjCNEioXzJobWead
O3dlRy85z-FBF7aw&sig2=Npmr3A_ryYyHqjXJKmlV5g
BRASIL. Ministério da Agricultura, Pecuária e Abastecimento. Regras para análise de
sementes. Ministério da Agricultura, Pecuária e Abastecimento. Secretaria de Defesa
Agropecuária. Brasília, DF: Mapa/ACS, 2009. 395p.
CARVALHO-PUPATTO, J. C.; BÜLL, L. T.; CRUSCIOL, C. A. C. Atributos químicos do
solo, crescimento radicular e produtividade de arroz de acordo com a aplicação de
escórias. Pesq. Agropec. Bras., v. 39, n. 12, p. 1213-1218, 2004.
http://www.scielo.br/pdf/pab/v39n12/22863.pdf
ENCICLOPÉDIA BIOSFERA, Centro Científico Conhecer - Goiânia, v.9, n.17; p. 1222
2013
COMISSÃO DE QUÍMICA E FERTILIDADE DO SOLO - RS/SC. Manual de adubação
e calagem para os Estados do Rio Grande do Sul e de Santa Catarina. 10.ed. Porto
Alegre, SBCS - Núcleo Regional Sul/UFRGS, 2004, 400 p.
CONN, E. E.; STUMPF, P. K. Introdução à bioquímica. São Paulo, Edgard Blücher,
1980, 525 p.
COPELAND, L. O.; McDONALD, M. B. Principles of seed science and technology.
4.ed. Boston, MA: Kluwer Academic Publishers, 2001, 467 p.
DATNOFF, L. E.; RODRIGUES, F. A.; SEEBOLD, K. W. Silicon and plant nutrition. In:
DATNOFF, L. E.; ELMER, W. H.; HUBER, D. M. (Eds.) Mineral nutrition and plant
disease. Saint Paul MN. APS Press, 2007, p. 233-246.
DATTA, S. K.; MUTHUKRISHNAN, S. Pathogenesis-related proteins in plants. Boca
Raton. CRC Press. 1999, 291 p.
DEREN, C. W.; DATNOFF, L. E.; SNYDER, G. H.; MARTIN, F. G. Silicon concentration,
disease response, and yield components of rice genotypes grown on flooded organic
histosols.
Crop
Science,
v.
34,
n.
3,
p.
733-737,
1994.
http://journals.cambridge.org/action/displayAbstract;jsessionid=1AC6B27F9A69E34267
E4DC75FDDE1C7E.journals?fromPage=online&aid=1287652.
https://www.crops.org/publications/cs/abstracts/34/3/CS0340030733?destination=public
ations%2Fcs%2Fabstracts%2F34%2F3%2FCS0340030733
FAO. World Agriculture: Towards 2015/2030. Summary Report. Corporate document
repository. 2002, 106p. http://www.fao.org/docrep/004/y3557e/y3557e00.htm
INTERNATIONAL SEED TESTING ASSOCIATION. ISTA. Handbook of variety
testing: electrophoresis testing. 1.ed, ISTA, 1992. http://seedtest.org/en/productdetail--0--0--0--38.html
JEBARA, S.; JEBARA, M.; LIMAM, F.; AOUANI, M. E. Changes in ascorbate
peroxidase, catalase, guaiacol peroxidase and superoxide dismutase activities in
common bean (Phaseolus vulgaris) nodules under salt stress. Journal of Plant
Physiology,
v.
162,
p.
929-936,
2005.
http://www.sciencedirect.com/science/article/pii/S0176161705000064
LIANG, Y. C.; MA, T. S.; LI, F. J.; FENG, Y. J. Silicon availability and response of rice and
wheat to silicon in calcareous soils. Comm. Soil Sci. Plant Anal., v. 25, p. 2285-2297,
1994. http://www.tandfonline.com/doi/abs/10.1080/00103629409369189#preview
LIMA FILHO, O. F. O silício e a resistência das plantas ao ataque de fungos
patogênicos. EMBRAPA – Empresa Brasileira de Pesquisa Agropecuária. Mato Grosso
do Sul, Artigo
1/8,
2005. Acesso
em
10/07/2012,
disponível
em:
http://www.cpao.embrapa.br/portal/artigos/artigos/artigo1.html#sdfootnote1sym
ENCICLOPÉDIA BIOSFERA, Centro Científico Conhecer - Goiânia, v.9, n.17; p. 1223
2013
LIMA FILHO, O. F. O silício é um fortificante e antiestressante natural para as plantas.
EMBRAPA – Empresa Brasileira de Pesquisa Agropecuária. Mato Grosso do Sul, Artigo
2/8,
2004.
Acesso
em
10/07/2012,
disponível
em:
http://www.cpao.embrapa.br/portal/artigos/artigos/artigo1.html#sdfootnote1sym
LIMA-FILHO, O. F.; TSAI, S. M. Crescimento e produção do trigo e da aveia branca
suplementados
com
silício.
EMPRESA
BRASILEIRA
DE
PESQUISA
AGROPECUÁRIA – EMBRAPA – Dourados, MS: Embrapa Agropecuária Oeste, 2007,
38 p. - Circular Técnico / Embrapa Agropecuária Oeste, ISSN 1679-0456; n. 41.
http://www.cpao.embrapa.br/publicacoes/online/zip/BP200741.pdf
MACHADO, A. A.; CONCEIÇÃO, A. R. Sistema de análise estatística para windows.
WinStat. Versão 2.0. Pelotas: UFPel, 2003.
MALONE, G.; ZIMMER, P. D.; CASTRO, M. A. S.; CARVALHO, I.; MENEGHELLO, G.
E.; PESKE, S. T. Identificação do estádio adequado para realização de análises
isoenzimáticas na caracterização de cultivares de arroz. Revista Brasileira de
Sementes,
v.
28,
n.
2,
p.
193-200,
2006.
http://www.scielo.br/pdf/rbs/v28n2/a25v28n2.pdf
MATAMOROS, M. A.; DALTON, D. A.; RAMOS, J.; CLEMENTE, M. R.; RUBIO, M. C.;
BECANA, M. Biochemistry and molecular biology of antioxidants in the Rhizobia-legume
symbiosis.
Plant
Physiology,
v.
133,
p.
499-509,
2003.
http://www.plantphysiol.org/content/133/2/499.full.pdf+html
MENEZES, S. M.; TILLMANN, M. A. A.; DODE, L. B.; VILLELA, F. A. Detecção de soja
geneticamente modificada tolerante ao glifosato por métodos baseados na atividade de
enzimas. Revista Brasileira de Sementes, v. 26, n. 2, p. 150-155, 2004.
http://www.scielo.br/scielo.php?script=sci_arttext&pid=S0101-31222004000200021
ORIOLI-JÚNIOR, V.; ARF, O.; COSTA, R. S.; BUZETTI, S. Modos de aplicação e doses
de silício em dois cultivares de trigo cultivados em semeadura direta. Scientia Agraria,
v. 9, n. 3, p. 377-383, 2008. http://redalyc.uaemex.mx/redalyc/pdf/995/99516777014.pdf
PERRY, C. C. Biosilicification – Structure, Regulation of Structure and Model
Studies, in Silicon Chemistry: From the Atom to Extended Systems (eds P. Jutzi and U.
Schubert), Wiley-VCH Verlag GmbH & Co. KGaA, Weinheim, Germany. doi:
10.1002/9783527610761.ch37,
2007.
http://onlinelibrary.wiley.com/doi/10.1002/9783527610761.ch37/summary
RAMIREZ, H.; CALDERON, A.; ROCCA, W. Técnicas moleculares para evaluar y
mejorar el germoplasma vegetal. In: ROCCA, W.; MROGINSKI, L. (Ed). Cultivo de
Tejidos en la Agricultura: Fundamentos y aplicaciones. Cali: CIAT, 1991, p. 825-856.
REIS, M. A.; ARF, O.; SILVA, M. G.; SÁ, M. E.; BUZETTI, S. Aplicação de silício em
arroz de terras altas irrigado por aspresão. Acta Sci. Agron., v. 30, n. 1, p. 37-43, 2008.
http://periodicos.uem.br/ojs/index.php/ActaSciAgron/article/view/1126/623
ENCICLOPÉDIA BIOSFERA, Centro Científico Conhecer - Goiânia, v.9, n.17; p. 1224
2013
SÁ, M. E. Importância da adubação na qualidade de sementes. In: SÁ M. E.; BUZZETI
S. (Ed.). Importância da adubação na qualidade dos produtos agrícolas. São Paulo:
Ícone, 1994, p. 65-98.
SANTOS, C. M. R.; MENEZES, N. L.; VILLELA, F. A. Alterações fisiológicas e
bioquímicas em sementes de feijão envelhecidas artificialmente. Revista Brasileira de
Sementes,
v.
26,
n.
1,
p.
110-119,
2004.
http://www.scielo.br/pdf/rbs/v26n1/a17v26n1.pdf
SANTOS, C. M. R.; MENEZES, N. L.; VILLELA, F. A. Modificações fisiológicas e
bioquímicas em sementes de feijão no armazenamento. Revista Brasileira de
Sementes, v. 27, n. 1, p. 104-114, 2005. http://www.scielo.br/pdf/rbs/v27n1/25187.pdf
SATTERS J. R. G.; ABDELGHANY, A.; ELBAGOURY, O.; WEST, S. H. Soybean seed
deterioration and response to priming: changes in specific enzyme activities in extracts
from dry and germinating seeds. Seed Science Research, v. 4, n. 1, p. 33-41, 1994.
SCANDALIOS, J. G. Genetic control of multiple molecular forms of enzymes in plants: a
review.
Biochemical
Genetics,
v.
3,
p.
37-39,
1969.
http://www.springerlink.com/content/m14r214544965557/
SINGH, S. P.; COOPER, J. E.; FISHER, H. L.; TARRANT, C. J.; LLOYD, T.; BANJO, J.;
CORFE, S.; JONES, C. Determining the chronology and components of psychosis
onset: The Nottingham Onset Schedule (NOS). Schizophrenia Research, n. 80, p. 117130, 2005. http://ac.els-cdn.com/S0920996405001696/1-s2.0-S0920996405001696main.pdf?_tid=5a02cfd217f32b577990848efa568886&acdnat=1339002828_b30799f61b
3da49867e6609cc552c0a9
SPINOLA, M. C. M.; CICERO, S. M.; MELO, M. Alterações bioquímicas e fisiológicas
em sementes de milho causadas pelo envelhecimento acelerado. Scientia Agrícola, v.
57,
n.
2,
p.
263-270,
2000. http://www.scielo.br/scielo.php?pid=S010390162000000200011&script=sci_arttext
STRECK, E. V.; KÄMPF, N.; DALMOLIN, R. S. D.; KLAMT, E.; NASCIMENTO, P. C. do;
SCHNEIDER, P.; GIASSON, E.; PINTO, L. F. S. Solos do Rio Grande do Sul. 2.ed.
rev. e ampl. Porto Alegre: Emater/RS, 2008, 222 p.
TOLEDO, M. Z.; GARCIA, R. A.; MERLINA, A; FERNANDES, D. M. Seed germination
and seedling development of white oat affected by silicon and phosphorus fertilization.
Scientia
Agricola,
v.
68,
n.
1,
p.
18-23,
2011.
http://www.scielo.br/pdf/sa/v68n1/v68n1a03.pdf
TUNES, L. M.; PEDROSO, D. C.; MENEGHELLO, G. E.; CASTRO, M. A. S.; BARROS,
A. C. S. A.; BADINELLI, P. G.; MUNIZ, M. F. B. Perfil enzimático em sementes de
ENCICLOPÉDIA BIOSFERA, Centro Científico Conhecer - Goiânia, v.9, n.17; p. 1225
2013
cevada em resposta a diferentes concentrações salinas. Interciência, v. 35, n. 5, 2010.
http://www.interciencia.org/v35_05/369.pdf
VIEIRA, A. R.; OLIVEIRA, J. A.; GUIMARÃES, R. M.; CARVALHO, M. L. M.; PEREIRA,
E. M.; CARVALHO, B. O. Qualidade de sementes de arroz irrigado produzidas com
diferentes doses de silício. Revista Brasileira de Sementes, v. 33, n. 3, p. 490-500,
2011. http://www.scielo.br/pdf/rbs/v33n3/12.pdf
VIEIRA, E. S. N.; VON PINHO, E. V. R.; CARVALHO, M. G. G.; SILVA, P. A.
Caracterização de cultivares de soja por descritores morfológicos e marcadores
bioquímicos de proteínas e isoenzimas. Revista Brasileira de Sementes, v. 31, p. 8694,
2009.
http://www.scielo.br/scielo.php?pid=S010131222009000100010&script=sci_arttext
VIGANO, J.; BRACCINI, A. L. E.; SCAPIM, C. A.; FRANCO, F. A.; SCHUSTER, I.;
MOTERLE, L. M.; TEXEIRA, L. R. Qualidade fisiológica de sementes de trigo em
resposta aos efeitos de anos e época de semeadura. Revista Brasileira de Sementes,
v. 32, n. 3, p. 86-96, 2010. http://www.scielo.br/pdf/rbs/v32n3/v32n3a10.pdf
YOSHIDA, S.; NAVESAR, S. A.; RAMIREZ, E. A. Effects of silica and nitrogen supply
one some leaf characters of rice plant. Plant and Soil, v. 31, p. 48-56, 1969.
http://www.springerlink.com/content/g745m3t081u83761/
ZAGO, A. S.; SILVA, C. A. T.; SILVA, T. R. B.; VIECELLI, C. A.; VERONA, E.; NOLLA, A.
Efeito de doses de silício no desenvolvimento da soja. Cultivando o Saber –
Cascavel,
v.
3,
n.
2,
p.
16-22,
2010.
http://www.fag.edu.br/graduacao/agronomia/csvolume32/03.pdf
ENCICLOPÉDIA BIOSFERA, Centro Científico Conhecer - Goiânia, v.9, n.17; p. 1226
2013